Cerebellar Outputs in Non-human Primates: An Anatomical Perspective Using Transsynaptic Tracers

Important insights into cerebellar function can be gained from an anatomical analysis of cerebellar output. Recent studies using transsynaptic tracers in nonhuman primates demonstrate that the output of the cerebellum targets multiple nonmotor areas in th

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25

Andreea C. Bostan and Peter L. Strick

Abstract

Important insights into cerebellar function can be gained from an anatomical analysis of cerebellar output. Recent studies using transsynaptic tracers in nonhuman primates demonstrate that the output of the cerebellum targets multiple nonmotor areas in the prefrontal and posterior parietal cortex, as well as the motor areas of the cerebral cortex. The projections to different neocortical areas originate from distinct output channels within the cerebellar nuclei. The neocortical area that is the main target of each output channel is a major source of input to the channel. Thus, a closed-loop circuit represents the fundamental macro-architectural unit of cerebro-cerebellar interactions. The outputs of these circuits provide the cerebellum with the anatomical substrate to influence the control of movement and cognition. Similarly, it has been shown that discrete multisynaptic loops connect the basal ganglia with motor and nonmotor areas of the cerebral cortex. Interactions between cerebro-cerebellar and cerebro-basal ganglia loops have been thought to occur mainly at the level of the neocortex. More recently, neuroanatomical studies demonstrate that the anatomical substrate exists for substantial interactions between the cerebellum

A.C. Bostan (*) Center for the Neural Basis of Cognition, Systems Neuroscience Institute and Department of Neurobiology, School of Medicine, University of Pittsburgh, Pittsburgh, PA, 15261, USA e-mail: [email protected] P.L. Strick Pittsburgh Veterans Affairs Medical Center, Pittsburgh, PA, 15261, USA and Center for the Neural Basis of Cognition, Systems Neuroscience Institute and Department of Neurobiology, School of Medicine, University of Pittsburgh, Pittsburgh, PA, 15261, USA e-mail: [email protected] M. Manto, D.L. Gruol, J.D. Schmahmann, N. Koibuchi, F. Rossi (eds.), 549 Handbook of the Cerebellum and Cerebellar Disorders, DOI 10.1007/978-94-007-1333-8_25, # Springer Science+Business Media Dordrecht 2013

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A.C. Bostan and P.L. Strick

and the basal ganglia in both the motor and nonmotor domains. These data, along with the revelations about cerebro-cerebellar circuitry, provide a new framework for exploring the contribution of the cerebellum to diverse aspects of behavior.

Introduction The neocortical areas that provide inputs to the cerebellum have been well established (Fig. 25.1) (Glickstein et al. 1985; Schmahmann 1996). On the other hand, the targets of cerebellar output are still in the process of being fully identified (Strick et al. 2009). Recent results from neuroanatomical studies using transsynaptic tracers in nonhuman primates indicate that cerebellar output targets both motor and nonmotor areas of the cerebral cortex. This feature of cerebellar output provides part of the neural substrate for the involvement of cerebellum not only in the generation and control of movement but also in nonmotor aspects of behavior. This chapter reviews new evidence about the areas of the cerebral cortex that are the target of cerebellar